2011 2nd International Conference on Environmental Science and Technology

IPCBEE vol.6 (2011) (2011) IACSIT Press, Singapore

Determination of Sexual Dimorphism in the Primary Wing and Tail Feathers of

A SUBSPECIES OF THE Rock Pigeon (Columba livia intermedia) using Principal
Component
Analysis, Elliptic Fourier Analysis and Discriminat Analysis
Queenilyn B. Albutra

Cesar G. Demayo

Department of Biological Sciences

College of Science and Mathematics
MSU-Iligan Institute of Technology
9200 Iligan City, Lanao del Norte, Philippines
queenilyn_albutra@yahoo.com

Department of Biological Sciences

College of Science and Mathematics
MSU-Iligan Institute of Technology
9200 Iligan City, Lanao del Norte, Philippines
c_gdemayo@yahoo.com

Mark Anthony J. Torres

Department of Biological Sciences
College of Science and Mathematics
MSU-Iligan Institute of Technology
9200 Iligan City, Lanao del Norte, Philippines
torres.markanthony@gmail.com
Absract Many studies conducted in the last decades have
shown that sexual dimorphism observed among species of
birds was largely influenced by social mating system and it is
more apparent in polygymous where male to male competition
frequent. However, studies conducted recently revealed that
there are other factors contributed to the presence of sexual
dimorphism. The primary aim of this study is to determine the
occurrence of sexual dimorphism in the monogamous species
of rock pigeon (Columba livia intermedia) based on the shape
variation of primary wing and tail feathers of the bird.
Analysis of shape variation was done using Principal
Component Analysis, Elliptic Fourier Analysis and
Discriminant Analysis. Results obtained shows that there is a
presence of sexual dimorphism in the species of rock pigeon
(Columba livia intermedia) and it is more apparent in the tail
feather than in the primary wing feather.Variation was related
to the preference of females over the males and not on male to
male competition and fight costs of tge rock pigeon.
Keywords-Columba livia intermedia; sexual dimorphism;
monogamous; Principal Component Analysis; Elliptic Fourier
Analysis; Discriminant Analysis

I.

INTRODUCTION

The Rock Pigeon (Columba livia), or Rock Dove, is a

member of the bird family Columbidae (doves and pigeons)
[1]. The species is generally monogamous, with two squabs
(young) per brood. Sexual dimorphism in birds is common
in nature and often attributable to sexual selection. This is
particularly true for sexual dimorphism in ornaments where
the exaggerated dimorphic traits are used predominantly in
the competition over mates [2]. Various studies were
conducted to determine the presence of sexual dimorphism

among the species of birds and often, the results were based
on the analysis of the birds body size and plumage
coloration.
Sexual dimorphism in body size in birds is commonly
regarded as a result of sexual selection favouring large
males through the size-advantage in combats over females
[3]. This idea is supported by the observation that
polygynous species frequently exhibit a larger degree of
sexual size dimorphism than do monogamous species [4];
although the pattern is far from perfect [5]. Models of
female choice, on the other hand, have concentrated on the
evolution of ornament size, such as tail length; since
polygynous species seem to possess more elaborate
ornaments than do monogamous species [6]. However, in
this study, another approach was made in establishing
sexual dimorphism in birds.
The primary aim of this study is to determine the
presence of sexual dimorphism in a subspecies of the rock
pigeon (Columba livia intermedia) based on the shape
variation of its primary wing and tail feathers. Examination
of shape variation was made using geometric
morphometrics such as Principal Component Analysis,
Elliptic Fourier Analysis and Discriminant Analysis. Also,
this study seeks to explain some underlying phenomena
involve in the evolution of the feathers that resulted in the
occurrence of sexual dimorphism within the species of rock
pigeon.
II. METHODOLOGY
During the last week of July 2010, a pair of rock pigeon
(Columba livia intermedia) species was collected to

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determine the presence of sexual dimorphism in the bird

species. In the laboratory, the primary wing feathers and the
tail feathers of the pigeon were taken carefully one by one.
The feathers were then scanned under 600 resolutions to get
a clear image of the feathers.
Using the tpsDIG program [7] and the bitmap images
of the scanned feathers, 100 closely spaced points were
made along the outline structure of each feather to obtain
the x and y coordinates. Mapping started at the most
proximal point where the barbs of the leading edge of the
feather met the rachis and ended at the analogous point on
the trailing edge of the feather. The tpsDIG program
facilitates the statistical analysis of landmark data in
morphometrics by making it easier to collect and maintain
landmark data from digitized images.
The landmark data was then subjected into a diffferent
multivariate analysis, which include Principal Component
Analysis (PCA), Elliptic Fourier Analysis (EFA) and
Discriminant Analysis, using the software PAST version
1.74 as platform[8].
III. RESULTS AND DISCUSSION
Following a series of analysis made in the primary
wing feathers and tail feathers of the rock pigeon (Columba
livia intermedia), it was found out that there was a shape
variation between the male and females feather of the said
bird species, consequently, suggesting the presence of
sexual dimorphism. Figure 1 shows the principal mean
shape of the feathers in each defined region of the rock
pigeon.

Figure1. Principal mean shape of the feathers in (A) females left primary
feather, (B) males left primary feather, (C) females right primary feather,
(D) males right primary feather, (E) females tail feather and (F) males
tail feather.

From the figure, it clearly illustrates that with respect to

its shape; the primary wing and tail feathers in male species
are quietly slender compared to that of females feathers.
Apart from this, there is no other significant variation
observed in the shape of the primary wing feather between
sexes. However, with regards to the mean shape of the tail
feathers in male and female rock pigeon. Another
distinction observed between the tail feathers was that, the
mean shape of the tail feathers in female species tend is
rounded in one end and tends to become pointed as it
approaches the other end while in the male species, the tail
feathers are more or less rounded on both ends. Aside from
variation in feathers shape, variation in the feathers size
was also evident. The mean shape of males tail feather is
longer compared to its female counterpart. As many
literatures would site, size variation is one important factor
that demonstrates sexual dimorphism among species of
birds.
The result of principal component analysis (PCA)
depicts that sexual dimorphism is more evident in the tail
feathers of the rock pigeon rather than in its primary wing
feathers. The principal component scores can be used as
observed values of morphological features in subsequent
analysis and in the case of this study, morphological
analysis of the feathers shape (Iwata and Ukai, 2002). Data
in table 1 implies that majority of the total variation shown
in the left and right primary wing feathers of the female
species was accounted only by the first principal component,
as shown in its % variance which is 95.39% and 95.18%
respectively. This would entail that the presence of variation
is explained only by 1 variable in the feather. However, the
case is different in the tail feathers of the rock pigeon. It
could be observed that the total variance in the data set does
not focus in only one principal component. Unlike in the
case of the primary feather, the majority of the total
variance is distributed in the three principal components
which have the highest % variance value. This would imply
that the shape variation in the tail feather was influenced
mainly by 3 variables present in the feather.
The same scenario was observed in the case of the male
species. The left and right primary wing feathers of the male
also shows the highest degree of variation with respect to its
first principal component while the variation in the tail
feathers were allocated in multiple principal components.
However, it can be observed that in male, apart from
the %variance value of the first three principal components,
the value of the fourth and fifth principal components are
relatively larger. These values were also quite larger
compared to the value of fourth and fifth principal
components in females tail feather. This would imply that
in males, the variation observed in the tail feathers was

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influenced greatly by more variables compared to that of

females.

TABLE1. % VARIANCE VALUE OF THE 5 PRINCIPAL COMPONENTS IN THE 3 DIFFERENT FEATHER REGIONS OF FEMALE AND MALE COLUMBA
LIVIA NTERNEDIA

Principal
Component
(PC)

1
2
3
4
5

Female
Left
95.392
1.8372
1.2626
0.87858
0.31897

Male

Right

Tail

Left

Right

Tail

95.183
2.6243
1.0358
0.54168
0.33042

50.111
31.121
12.451
3.1254
2.0745

93.195
4.0163
1.1637
0.94147
0.38226

94.238
3.872
0.82487
0.50664
0.25245

36.082
30.328
16.868
8.452
5.3013

Discriminant analysis is another tool used in this study

that supplements the results obtained in the principal
component analysis for the presence of sexual dimorphism
in rock pigeon based on its feathers shape. The visual result
of this analysis was shown in a form of histogram as shown
in figure 2.
The histogram shows the difference between the shape
of male and females feather in each of the defined region.

Based on the graphs, it clearly shows that sexual

dimorphism is evident in all regions of the birds since there
is no presence of overlaps in the graph. But it is in the tail
region of the rock pigeon that exhibits the greatest sexual
dimorphism among them all due to the existence of large
gap between the two components. The farther the
components are from each other, the greater is the
difference in the shape of the feathers.

Figure2. Histogram showing the variation in the feathers shape of the male (blue) and female (red) species of Columba livia intermedia between (A) the left
primary wing region, (B) the right primary wing region and (C) the tail region.

Conversely, there were instances wherein sexual

dimorphism is still apparent even if overlapping occurs in
the histogram. This can be explained by the values yielded
in the discrimination score table where the percentage of
correctly classified items, as shown in table 3, was given.
This percentage shows how correctly female feathers were
classified as female and how correctly male feathers were
classified as male. If discriminant analysis is effective for a
set of data, the classification table of correct and incorrect
estimates will yield a high percentage correct. In the given

set of data, the result shows that in the primary left wing
region and tail region of the bird, 100% of the male and
female feathers were classified correctly. In the primary
right wing region, on the other hand, only 97.5% of the
male and female feathers were classified correctly. One
male feather was mistakenly classified as female feather.
But, despite this discrepancy, it is still safe to say that the
degree of shape variation in the right primary wing feather
is relatively high since the minimum percentage of correctly

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classified items should be 75% for it to be considered

significant.

TABLE 2. PERCENTAGE OF PERFECTLY CORRECTED FEMALE AND MALE FEATHERS IN THE 3 DIFFERENT REGIONS

Left Primary Feathers

Right Primary Feathers

Tail Feathers

Female

Male

% Corrected

Female

Male

% Corrected

Female

Male

% Corrected

Female

20

100%

20

100%

12

100%

Male

20

100%

19

95%

12

100%

Total

100%

97.5%

Many studies were conducted with regards to the

sexual dimorphism in the birds and most frequently; these
studies utilized the body size and ornaments of the birds. In
a study conducted to determine which morphological
measurements was more efficient in establishing sexual
dimorphism, result showed that the degree dimorphism was
highest for bill depth measurements and lowest for feather
measurements (wing and tail lengths) [9]. However, in this
study, only the feathers were taken into consideration so the
discussion would focus mainly on factor or factors that
might cause the variation of the feathers especially in the
tail that in turn, determines the sex of the birds. Nonetheless,
it could also be consider that the variation of the feathers
size might as well correlate on the variation of the feathers
shape.
In 1871, Charles Darwin argued that sexual
dimorphism evolved by means of sexual selection which
has two distinct types [10]. The first type of sexual selection
is the intrasexual selection wherein individuals of one sex,
typically the males, evolve traits that enable them to
compete with other individuals of the same sex and win
mating opportunities. The other type is the intersexual
selection, wherein the individual of one sex evolve traits
that are preferred by the members of the opposite sex,
typically the females. This is referred to as mate choice. In
Darwins sexual selection theory, it was pointed out that
sexual dimorphism is dependent on the reproductive
structure of the population (monogamy, polygyny, panmixia)
and that sexual dimorphism for size is more often found and
more pronounced in polygynic species than in monogamic
ones. Thus, intrasexual selection is more apparent than
intersexual selection.
In this study, however, the presence of evident sexual
dimorphism in the tail feathers of rock pigeon (Columba
livia intermedia) is related to intersexual selection. This
statement is due to the fact that the rock pigeon is an
example of monogamous species, wherein single female
species would only mate with single male species.
Therefore, there is no need for the males to develop a
favourable trait in order to win mating opportunities since
no male to male competition is involved. In this case, the
evolution of a narrower and elongated tail feathers in rock
pigeon has something to do with the females preference
over the males. Females may choose males that appear to

100%

be strong and healthy, thus likely to possess good alleles

and give rise to healthy offspring [11]. However, in some
species females seem to choose males with traits that do not
improve offspring survival rates, and even traits that reduce
it, potentially leading to traits like the peacock's tail [12].
The results obtained in this study is consistent with the
earlier study conducted in female barn swallows (Hirundo
rustica)[13,14,15]. In the study, the male tail ornaments are
the target of a strong directional female mate preference.
Further, it was suggested that used asymmetry in secondary
sexual characters to determine the males quality, but now it
has shown that female barn swallows prefer males with
symmetric tails. Within the species, males with longer tails
have more symmetric tails, so tail length may reliably
reflect the males quality as predicted by good-gene models.
These studies conducted are consistent with the results
obtained in this study since rock pigeons also possess
symmetrical tail feathers with more or less having the same
length and resembles a fan when spread out. If the results of
the studies on swallows would be used as basis in the
analysis of the current study, most likely, this subspecies of
female rock pigeon would prefer males with longer and
symmetrical tails for it reflects a males ability to possess a
good quality of alleles which insures a strong and healthy
offspring.
In another study that relates the tail dimorphism to the
flight cost of the bird using a comparative analysis and
aerodynamic theory, it was shown that there are costs of tail
elongation for different species of birds [16]. The result of
the study shows that natural selection, in a form of flight
costs, plays an important role in determining the
phylogenetic distribution of tail type and length. The
analysis is based on a recent model of lifting surfaces which
applies classic aerodynamic studies of airplane wings to the
lift and drag associated with birds tail. Species which rely
heavily on flight should have less aerodynamically costly
tails, which means that the ratio of dragged tail over lifted
tail should be small. Results also provide evidence of a link
between the strength of sexual selection and tail types. In
species with tails that have a large aerodynamic cost, there
is large sexual dimorphism in tails length.
As mentioned earlier, a spubspecies of the rock pigeon
(Columba livia intermedia) is a non-migratory bird that
does not rely mostly on flights and therefore possesses a

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graduated type of tail. Species which acquire this type of

tail have high aerodynamically costly tail. Thus, this would
imply that in this subspecies of rock pigeon, its tail is
mostly dragged on the ground than lifted in the air. This
might be the reason why sexual dimorphism in the rock
pigeon is more evident in the tail region than in the primary
wing region. The presence of sexual dimorphism in the tail
of rock pigeon is supported by the idea that sexual
dimorphism is evident in species of birds having large
aerodynamically costly tails [16].
Based on the supporting studies given above, there was
no direct explanation provided to elucidate the phenomena
of sexual dimorphism based on feathers shape. All cited
cases of sexual dimorphism in birds were primarily based
on the size variation in tails feather, which is also evident
in this study. Thus, it could be assume that there might be a
correlation between the shape variation and size variation in
the feathers of rock pigeon (Columba livia intermedia),
especially in the tail region. It could be speculated that as
the size of the feather differentiate as a result of intersexual
selection and natural selection, its shape also follows; hence,
resulting in the presence of sexual dimorphism in the tail
feathers of rock pigeon (Columba livia intermedia) with
respect to its size and shape.

[3]

[4]
[5]
[6]
[7]
[8]

[9]

[10]
[11]
[12]
[13]

[14]

IV. CONCLUSION

[15]

From the results obtained in this study, it is concluded

that sexual dimorphism is apparently present in the
monogamous subspecies of the rock pigeon (Columba livia
intermedia). Shape variation in the feathers between male
and female species is the major basis for this finding.
Sexual dimorphism is evident in both primary wing feathers
and tail feathers of the bird species; however, it is more
evident in the tail region. Male species have narrower and
longer tail feathers compared to that of females. Factors that
contribute to the feathers variation include intersexual
selection and natural selection. However, these two factors
did not directly elucidate the relationship between shape
variation and sexual dimorphism. Intersexual selection and
natural selection only explicate the presence of sexual
dimorphism in birds with respect to the feathers length.
Thus, in this study, it was just speculated that since size
variation in the tail feather is also evident, there might be a
correlation between the size and shape variation in the
feathers. Further studies are still needed in order to verify
this assumption.

[16]

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